Some Thermal Requirements of Fiddler Crabs of the Temperate and Tropical Zones and Their Influence on Geographic Distribution

Thermal stress is one environmental parameter that has greatlyinfluenced the migration of crustaceans from the sea to land.Since a greater number of species of terrestrial crabs are foundin the tropics than in the temperate zone, comparative studiesof the influence of temperature on latitudinally separated populationswere undertaken. Two tropical species, U. rapax and U. thayeri,may occur as far north as St. Augustine, Florida, or, followinga severe winter, may be rare north of Cape Kennedy. The lethaleffect of the low temperatures recorded during one severe winter(1957–58) is supported by laboratory studies in whichLD₅₀, deaths occurred in 4.5 days at 10°C for U. rapax acclimatedto 18°C. The experiment demonstrates that U. rapax cannotacclimate to and survive low temperatures. This contrasts markedlywith the situation in semi-terrestrial crabs of the temperatezone, which are able to acclimate to cold. The distribution of Uca around Cape Cod Bay correlates wellwith the coastal hydrographic thermal gradient and supportsPassano's suggestion that temperatures below 20° may belimiting as they inhibit proecdysis in U. pugnax. Such an inhibitionis found experimentally in U. pugilator and in the tropicalspecies, U. rapax. It is hypothesized that a shift in the thermodynamicsof the processes underlying molting has not occurred in Ucaof the temperate zone. The paucity of semi-terrestrial Brachyurain the temperate zone may be due to the failure of many speciesto evolve capacity-adaptations to carry out all requisite lifeprocesses at temperatures below 20°, or the resistance-adaptationsnecessary to survive the low temperatures of winter.     

Causes and consequences of thermal tolerance limits in rocky intertidal porcelain crabs, genus petrolisthes

Vertical zonation of intertidal organisms, from the shallowsubtidal to the supralittoral zones, is a ubiquitous featureof temperate and tropical rocky shores. Organisms that livehigher on the shore experience larger daily and seasonal fluctuationsin microhabitat conditions, due to their greater exposure toterrestrial conditions during emersion. Comparative analysesof the adaptive linkage between physiological tolerance limitsand vertical distribution are the most powerful when the studyspecies are closely related and occur in discrete vertical zonesthroughout the intertidal range. Here, I summarize work on thephysiological tolerance limits of rocky intertidal zone porcelaincrab species of the genus Petrolisthes to emersion-related heatstress. In the eastern Pacific, Petrolisthes species live throughouttemperate and tropical regions, and are found in discrete verticalintertidal zones in each region. Whole organism thermal tolerancelimits of Petrolisthes species, and thermal limits of heartand nerve function reflect microhabitat conditions. Speciesliving higher in the intertidal zone are more eurythermal thanlow-intertidal congeners, tropical species have the highestthermal limits, and the differences in thermal tolerance betweenlow- and high-intertidal species is greatest for temperate crabs.Acclimation of thermal limits of high-intertidal species isrestricted as compared to low-intertidal species. Thus, becausethermal limits of high-intertidal species are near current habitattemperature maxima, global warming could most strongly impactintertidal species.     

Adaptive significance of hatching rhythms and dispersal patterns of estuarine crab larvae: avoidance of physiological stress by larval export?

Estuarine crabs commonly display two larval dispersal patterns in which larvae are either exported from or retained within estuaries. The semiterrestrial fiddler crab Uca minax (LeConte, 1855) hatches on nocturnal spring high tides in the upper estuary and larvae are rapidly transported downstream. The mud crab Rhithropanopeus harrisii (Gould, 1841) hatches on nocturnal high tides of any amplitude and larvae are retained behaviorally in the upper estuary throughout development. If larvae are exported from the estuary to avoid environmental stress, then exported larvae should be less tolerant of high temperatures and low salinities than retained larvae. Larvae of these two species of estuarine crabs were hatched at 20‰ and 25 °C and subjected to salinities of 0, 5, 10,20, and 30‰, temperatures of 25 and 35 °C, and exposure times of 2, 6, 12, and 48 h. Larvae of both species reared at 30 or 20‰ survived well, while those reared in fresh water all died within 2 h regardless of temperature. Mud crab larvae reared at 5 and 10‰ survived better at the lower temperature (25 °C), higher salinity, and shorter exposure times. There was no significant effect of temperature or salinity on the survival of fiddler crab larvae, although survival decreased with increasing exposure time. Thus, the hypothesis that fiddler crab larvae are exported into stable coastal waters to reduce physiological stress is not supported. However, fiddler crab larvae may have evolved to be very tolerant of extreme temperature and salinity stress because they, unlike mud crabs, often release their larvae into shallow creeks. Most fiddler crab larvae are released on nocturnal spring high tides, which facilitates dispersal from tidal creeks. However, freshwater runoff and heat transferred from the marsh surface to flooding waters may still create stressful conditions for larvae soon after they are released. Larval release on spring high tides may facilitate dispersal from tidal creeks.     

Stimulation of feeding behavior in three species of fiddler crabs by hexose sugars

Feeding responses to five hexoses were examined in three closelyrelated species of fiddler crabs, Uca minax, U.pugnax and U.pugilator.Hexoses tested were glucose, galactose, sorbose, fructose andmannose. Intact crabs and eyestalk-ablated crabs were tested.Responses to sugars were species specific. Eyestalks are directlyinvolved in vision and overall neural integration as well aswith chemosensory and metabolic pathways associated with feeding.Overall, eyestalk-ablated crabs were more sensitive uian intactcrabs. Studies of responses of individuals within a populationto hexoses showed tiiere are individual U.pugnax that respondto galactose and others that do not. Similarly, there were U.pugilatorindividuals that were mannose responsive and others that didnot respond to mannose. An additional study of differences inpopulation responses to hexoses would provide valuable toolsin studying geographic relationships between fiddler crab populations.     

Comparative Studies of Reproductive Behavior in Mantis Shrimps and Fiddler Crabs

Comparative ethological studies of closely related species canplace into clear perspective the functions of behavioral traitsas species diverge and adapt to different environments. In thispaper we contrast the reproductive behavior of species withintwo crustacean groups: the mantis shrimps (stomatopods) andthe fiddler crabs (genus Uca). For the stomatopods, we identifyprobable selective relationships between features of species'ecology and their mating systems. Population density, the kindand availability of shelters in which these shrimps live, theintensity of predation and spawning cycles all play importantroles in molding reproductive behavior in this group. Associationsbetween the ecology and mating systems of fiddler crabs havebeen discussed recently elsewhere. Here we focus on a comparativeanalysis of sexual communication in these crabs. A study ofcompetitive courtship signaling and mate choice in the fiddlercrab Uca beebei has shown that certain male courtship signalsare highly attractive probably because they exploit female sensory-responsesystems that have been molded by selection for escape from predators.Interspecific comparison of male courtship displays and theresponses of females to these displays suggest that sensoryexploitation may play an important role in the evolution ofsexual signals in the genus. Comparative studies have advancedour understanding of how natural and sexual selection affectthe reproductive behavior of both stomatopods and fiddler crabs.     

Predation by red-jointed fiddler crabs on congeners: interaction between body size and positive allometry of the sexually selected claw

The enlarged (major) claw of male fiddler crabs is used in contestsover breeding burrows and is waved to attract females. We recentlydiscovered that males of the red-jointed fiddler crab, Uca minax,also use the claw to kill smaller-sized fiddler crabs, U. pugnaxand U. pugilator, with which they co-occur in Atlantic coastsalt marshes. Large U. minax males use walking legs or the enlargedclaw to capture prey feeding on moist sand flats. On sand flats,small U. minax males and females are much less common than largemales, suggesting that large males move onto sand flats to seekprey. Males of prey species use the major claw against attackingpredators and, consequently, are more likely than females toescape. In laboratory experiments, large U. minax males weremore likely to attack and kill small-clawed males and femalesthan large-clawed males, consistent with a preference for morevulnerable, less threatening prey. The size of the major clawis a positive allometric function of body size. The allometricfunction varies little among species. Also, the mechanical advantageand indices of closing speed and closing force of the majorclaw, when corrected for body size, are not consistently greaterin U. minax relative to prey species. Thus, predation by U.minax males may reflect the opportunity afforded by larger bodysize and positive allometric growth, which result in a majorclaw that is more massive than the prey it is directed against.     

Habitat partitioning between prey soldier crab Mictyris brevidactylus and predator fiddler crab Uca perplexa

Interaction and habitat partition between the soldier crab Mictyris brevidactylus (prey) and the fiddler crab Uca perplexa (predator) were examined at a sandy tidal flat on Okinawa Island, Japan, where they co-occur. Both live in dense colonies. When the soldier crabs were released in the densely populated habitat of the fiddler crab, male fiddler crabs, which maintain permanent burrows in hard sediment, preyed on small soldier crabs and repelled large ones. Thus, the fiddler crabs prevented the soldier crabs from trespassing. It was also observed whether soldier crabs burrowed successfully when they were released 1) where soldier crab burrows just under the sand were abundant, 2) in a transition area between the two species, 3) an area without either species, and 4) where artificial tunnels simulated soldier crabs' feeding tunnels were made by piling up sand in the area lacking either species. In contrast to the non-habitat area, many soldier crabs burrowed in the sediment near the release point in the tunnel, transition and artificial tunnel areas. This indicates that the feeding tunnels on the surface attracted other crabs after emergence. When the large male fiddler crabs were transplanted into the artificial burrows made in soft sediment of the soldier crab habitat, all left their artificial burrows by 2 days. In the fiddler crab habitat, however, about one-third of the transplanted male fiddler crabs remained in the artificial burrows after 3 days. The soldier crabs regularly disturb the sediment by the up and down movement of their burrow (small air chamber) between tides. This disturbance probably prevents the fiddler crab from making and occupying permanent burrows. Thus, it appears that these crabs divide the sandy intertidal zone by sediment hardness and exclude each other by different means.     

Temperature influences the reproduction of fiddler crabs at the southern edge of their distribution

Understanding the spatial patterns of reproductive biology might provide predictions about fitness and population stability in different locations within the geographical range of a particular species. Leptuca uruguayensis is a fiddler crab that breeds year round in tropical estuaries but only in summer months in temperate salt marshes. In this study, we examined several reproductive attributes of the southernmost population of L. uruguayensis, including the proportion of ovigerous females, the proportion of surface‐active crabs, the developmental status of the gonads and hepatopancreas, and the fullness of seminal receptacles, and related them to environmental factors such as temperature, photoperiod, and sediment organic matter content. We found that temperature was the environmental factor that was most correlated with the reproductive process of the southernmost fiddler crab, since this environmental factor was related to ovarian development, to the fullness of seminal receptacles, and to the hepatosomatic index. At the southern edge of its distribution, the low temperatures of winter restricted reproduction in L. uruguayensis. These winter temperatures might represent the lower limit of the thermal window of this fiddler crab, limiting its extension toward higher latitudes.     

Habitat and host specificity of trematode metacercariae in fiddler crabs from mangrove habitats in Florida

Fiddler crabs (Uca spp.) are common inhabitants of temperate and tropical coastal communities throughout the world, often occupying specific microenvironments within mangrove and salt marsh habitats. As second intermediate hosts for trematodes, we investigated patterns of host distribution and parasitism for 3 species of sympatric fiddler crabs in mangrove habitats adjacent to the Indian River Lagoon, Florida. Fiddler crab distribution varied among species, with Uca speciosa dominating the low and mid intertidal regions of mangrove banks. This species also exhibited higher prevalence and abundance of Probolocoryphe lanceolata metacercariae compared with Uca rapax, which is relatively more abundant in the high intertidal zone. We conducted a field experiment to test whether U. speciosa was more heavily parasitized by P. lanceolata as a result of its habitat distribution by raising U. speciosa and U. rapax under identical environmental conditions. After exposure to shedding cercariae under the same field conditions, all individuals of U. speciosa became parasitized by P. lanceolata, whereas no U. rapax were parasitized, suggesting that differences in parasitism were driven by host selection.     

Timing of hatching and release of larvae by brachyuran crabs: patterns, adaptive significance and control

Most semiterrestrial, intertidal and shallow subtidal brachyuran crabs that live in tropical and warm temperate estuaries, bays and protected coasts world-wide release their planktonic larvae near the times of nocturnal high tides on the larger amplitude tides in the biweekly or monthly cycles of tidal amplitude. Crab larvae usually emigrate quickly to the sea where they develop to return as postlarvae to settle in habitats suitable for their survival. Predators of larvae are more abundant where larvae are released than where they develop, suggesting that this migration from estuaries to the sea reduces predation on larvae. Crabs with larvae that are relatively well-protected by spines and cryptic colors do not emigrate and often lack strong reproductive cycles, lending support to this explanation. Adults control the timing of the release of larvae with respect to the biweekly and monthly cycles of tidal amplitude by controlling when they court and mate and females control when development begins by controlling when they ovulate and allow their eggs to be fertilized by stored sperm. By changing the time they breed, fiddler crabs (Uca terpsichores) compensate for the effects of spatial and temporal variation in incubation temperature on development rates so that embryos are ready to hatch at the appropriate time. Control of the diel and tidal timing of hatching and of release of larvae varies with where adults live. Females of the more terrestrial species often move from protected incubation sites, sometimes far from water, and they largely control the precise time, both, of hatching and of release of larvae. Females of intertidal species also may influence when embryos begin to hatch. Upon hatching, a chemical cue is released that stimulates the female to pump her abdomen, causing rapid hatching and release of all larvae in her clutch. Embryos, rather than females, largely control hatching in subtidal species, perhaps because females incubate their eggs where they release their larvae. Topics for further study include the mechanism whereby adults regulate the timing of breeding, the mechanisms by which females control development rates of embryos, the nature of communication between females and embryos that leads to precise and synchronous hatching by the number (often thousands) of embryos in a clutch, and the causes of selection for such precision. The timing of hatching and of release of larvae by cold-temperate, Arctic, and Antarctic species and by fully terrestrial and freshwater tropical species has received little attention.     

The Effect of Low Temperature upon Starch, Sucrose and Fructan Synthesis in Leaves

Measurements on plants in a temperature gradient tunnel showthat diurnal accumulation of starch in illuminated leaves wasgreatly reduced at temperatures below 8 °C, whereas sucrosesynthesis was less affected under similar conditions. High chillingsensitivity for leaf starch accumulation was observed in a numberof chilling resistant temperate species. Enzymes of sucroseand fructan metabolism from mature leaves of Lolium temulentumwere less strongly inhibited at low temperatures than enzymesinvolved in starch synthesis. These results are discussed inrelation to carbon partitioning in species which grow and aremetabolically active at chilling temperatures. Lolium temulentum, starch, sucrose, fructan, temperature, enzyme activity, carbon partitioning     

Shell use by juvenile fiddler crabs Uca pugnax and U. pugilator

The present study documents for the first time shell use by juvenile fiddler crabs in the salt marsh. Twenty visits were made to six salt marsh sites at Tybee Island, Georgia between 2007 and 2009. One hundred empty Littorina irrorata shells were collected at each site on each field trip. Juvenile carapace width was measured, crabs sexed, and species identification completed using RFLP analysis. Shell use of up to 79% was observed. Two species of fiddler crabs were found in empty shells, Uca pugnax and U. pugilator. U. pugnax was the dominant species at all sites representing 62-84% of the juvenile fiddler crab population. Juvenile sex ratios were female-biased (1.7:1) at all six sites. Juvenile size did not vary significantly between species but males of both species were significantly larger than females. Size frequency distribution of carapace width revealed that shell use varied with size and sex. In the 3 to 4 mm size class, juvenile females outnumbered juvenile males in empty L. irrorata shells while in the 5 to 6 mm size class and greater, juvenile males outnumbered juvenile females in shells. Significantly more juvenile fiddler crabs were found in empty shells during flood than ebb tide at 3 of the sites. This discovery illuminates the resourcefulness of juvenile fiddler crabs and provides another mechanism that might enhance survival.     

Visual and Acoustical Signalling during Courtship by Fiddler Crabs (Genus Uca)

Male fiddler crabs (Genus Uca) employ both visual and acousticalsignals to attract females for mating. In U. pugilator and severalother American species, the males attract females during theday first by waving, then by producing sounds just within theirburrows. At night, the males produce sounds at low rates, butwhen touched by a female, they increase their rate of soundproduction. In the European species, U. tangeri, many elements of courtshipare similar to those in U. pugilator, but two types of soundsare produced. One of these, the short drumwhirl, appears tosubstitute for waving when the male is temporarily obscuredfrom the female during his diurnal courtship activities. Thelong drumwhirl is used under different circumstances. The acoustical responses of a male to a female influence thecourtship behavior of other males in the area. When sounds fromstimulated males are played back to test males during the day,their lates of waving increase. At night, the playbacks elicitincreases in rates of sound production. The influence of tidal oscillations, temperature, and lightcycles on the behavior of males is discussed. Courtship activities of aquatic crabs are compared to thoseof terrestrial Brachyura. In aquatic forms, courtship may beabsent or, if present, does not involve elaborate signallingby the male. Chemical or visual cues at close range are themost important stimuli. In several genera of terrestrial crabs,visual signalling for prolonged periods is common, and soundsare often emitted by males to "call" females from their burrowsto the surface for mating. Some of the factors that may accountfor differences in courtship activities in aquatic and terrestrialspecies are discussed.     

Reproducing on Time When Temperature Varies: Shifts in the Timing of Courtship by Fiddler Crabs

Many species reproduce when conditions are most favorable for the survival of young. Numerous intertidal fish and invertebrates release eggs or larvae during semilunar, large amplitude, nocturnal tides when these early life stages are best able to escape predation by fish that feed near the shore during the day. Remarkably, some species, including the fiddler crabs Uca terpsichores and Uca deichmanni, maintain this timing throughout the year as temperature, and thus the rate of embryonic development, vary. The mechanisms that allow such precision in the timing of the production of young are poorly known. A preliminary study suggested that when temperature decreases, U. terpsichores mate earlier in the tidal amplitude cycle such that larvae are released at the appropriate time. We tested this idea by studying the timing of courtship in U. terpsichores and U. deichmanni as temperature varied annually during two years, at 5 locations that differed in the temperature of the sediment where females incubate their eggs. Uca terpsichores courted earlier at locations where sediment temperature declined seasonally but not where sediment temperature remained elevated throughout the year. In contrast, clear shifts in courtship timing were not observed for U. deichmanni despite variation in sediment temperature. We discuss other mechanisms by which this species may maintain reproductive timing. These two species are likely to be affected differently by changes in the frequency and intensity of cold periods that are expected to accompany climate change.     

The effect of abrupt salinity change on oxygen consumption in crab megalopae of Chincoteague Bay,USA

Variability in salinity is an environmental stressor that crab megalopae encounter as they are carried by tides and currents throughout Chincoteague Bay. We exposed blue crab (Callinectes sapidus) and fiddler crab (Uca spp.) megalopae to abrupt salinity changes from 10 to 31 ppt and measured their oxygen usage. It was hypothesized that the megalopae would cope with the changes in a manner reflective of the documented abilities and tolerances of adult crabs. It was also hypothesized that lower salinities would have a particularly detrimental effect on the megalopae reflected by both increased oxygen usage and mortality. The megalopae of both species did exhibit an increase in oxygen use at lower salinities, although the effect was more pronounced during the initial transition and decreased during acclimation. The megalopae mirrored the adult responses, with blue crab larvae consuming more oxygen per mg of wet weight at lower salinities, whereas fiddler crab larval oxygen consumption was relatively uniform at all salinities. Mortality of some blue crab postlarvae was observed at 10 ppt while all larval fiddler crabs survived. Coupled with the introduction of additional fresh water into the global water system, these results indicate that further investigation into this subject is necessary.     

Nouvelles formulations des regles ecologiques connues sous le nom de reegle de Bergmann et loi de Jordan: Amendments of the ecological rules known as Bergmann's and Jordan's rules

Measurements of populations of planktonic and micronektonicspecies from the Atlanto-mediterranean region (about 10,000specimens) have lead to improvements in Bergmann's rule relatingto the increase in size with latitude. There are three cases : Mediterranean specimens may be smaller(e.g. the mysid Eucopia hansent), equal to (e.g. the decapodcrustacean Gennadas elegans) or larger than Atlantic ones (e.g.the thecosomatous genus Cymbulia). All the species in the last group have their principal distributionin warm ocean waters. In the Atlantic, tropical species (e.g.the euphausiid Euphausia gibboides) decrease in size in thenorthern parts of their range while temperate species (e.g.the decapod crustacean Sergesles corniculum) decrease in sizetowards both the pole and the equator. The comparison with the number of vertebrae of fish, which increasesas does the size according to Jordan's rule, is interesting:though correct for cold water species (e.g. Merlucdus merluccius),this rule cannot be applied to warm water species (e.g. Merlucciussenegalensis). Thus, these rules must be corrected like this:- "a species reachesits maximal size in high, intermediate or low latitudes of itsarea if it has a boreal, temperate or tropical distribution"; - "the number of vertebrae increases with latitude for borealfish and decreases for tropical ones".     

Thermal tolerance of tropical and temperate alpine plants suggests that ‘mountain passes are not higher in the tropics’

Aim

Tolerance of species to extreme temperatures largely determines their distribution and vulnerability to climate change. We examined thermal tolerance in tropical and temperate alpine plants, testing the hypotheses that: (a) temperate plants are resistant to more extreme temperatures and have an overall wider thermal tolerance breadth (TTB); (b) TTB in temperate plants is wider than TTB in tropical plants during the entire growing season; (c) resistance to frost and heat varies during the season in temperate plants but not in tropical plants; (d) TTB of a species predicts its latitudinal range.

Location

Tropical (Ecuador, Bolivia) and temperate (USA, Austria) mountains.

Time period

Four periods of the growing season (2014, 2016–2019).

Major taxa

Ninety-six vascular plant species.

Methods

We employed the electrolyte leakage method to estimate the temperature resistance, that is, the temperature at which 50% tissue injury (Lt50) occurs in leaves. We used phylogenetic linear mixed-effect models in a Bayesian framework to test for differences between the plant groups.

Results

Temperate and tropical plants do not differ in their temperature resistance. The four hypotheses are rejected since: (a) temperate plants do not have significantly wider overall TTB compared to tropical plants, (b) TTB of temperate plants is wider than TTB of tropical plants only at the end of the temperate summer, (c) seasonal acclimation is observed in both plant groups, (d) the latitudinal range of the plants is not related to TTB.

Main conclusions

The lack of TTB differences between temperate and tropical alpine plants is consistent with trends observed in ectothermic animals, which suggests a general latitudinal pattern in high-elevation poikilotherm organisms. Limited acclimation capacity to cope with long freezing exposures restricts the occurrence of tropical alpine species to thermally aseasonal environments making them particularly vulnerable to climate change.     

Life-history strategies of Australian lizards: a comparison between the tropics and the temperate zone

Summary Previous studies have suggested that tropical and temperate-zone lizards may differ fundamentally in life histories. We tested the applicability of this idea to Australian species by comparing temperate-zone species of agamid and scincid lizards with their congeners from the seasonal tropics. Data were derived from dissection of 1,941 specimens and from published information. Clutch size and egg size were positively correlated with mean maternal body size in most lizard species from both climatic zones. Mean body size of the lizards studies did not differ between the tropics and the temperate zone, nor did egg or hatchling size. However, tropical skinks showed considerably (approximately 20%) lower clutch size and relative clutch mass than did temperate-zone skinks. This difference was partly due to the higher incidence of species with low, invariant clutch size in the tropical lizard fauna (as seen in other continents as well), but primarily due to a trend for lineages (especially genera) with relatively high fecundity to be more common in the temperate zone than in the tropics. In contrast to studies on African lizards, our data suggested that modification of clutch size between areas has not occurred within genera: congeneric species from the tropics and temperate zone did not differ in clutch size. Production of more than one clutch per annum by individual females was common in both climatic zones. Tropical lizards may differ from temperate-zone species in showing higher reproductive frequencies, more rapid growth and earlier maturation. However, most of these effects may be due to phenotypic responses to environmental conditions (especially longer annual activity season), rather than to genetically based lifehistory adaptations.